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A higher-order theory of emotional consciousness

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This means that the inclusion of fearful emotions in the preceding movie had a clear effect on the level of subsequent resting state activity. The stronger resting state effects of the preceding emotional movies are further confirmed by the more delayed recovery of the signal changes during the resting state period 90 s after emotional movies. This study clearly demonstrates that emotions have an impact on the subsequent resting state thus implying what we coined as stimulus-rest interaction see Northoff et al. This was also observed in another study.

This revealed increased functional connectivity from the amygdala to the cortical midline structures like the MPFC, the PCC, and the precuneus. This indicates that psychological stress implicating strong and negative emotions can affect the subsequent resting state activity thus implying stimulus-rest interaction.

Taken together, these studies demonstrate the close relationship between resting state activity and emotion-related activity. This seems to be especially apparent in the midline regions as core nucleus of the DMN. The high intrinsic activity in these regions seems to be closely related to emotion processing in though yet unclear ways. Different emotions seem to modulate the degree of stimulus-related deviation from the high resting state activity in different ways. The close relationship between emotions and resting state is further supported albeit indirectly by the observation of severe resting state alterations in major depressive disorder see Alcaro et al.

For that I turn to two of the major proponents of emotional feeling, Jaak Panksepp and Jim Russell, and how they conceptualize especially the subjective-experiential component of emotional feeling. This will be then linked in subsequent sections to the above described findings of the close relationship between intrinsic activity and emotions. Based on the centrality of affect and emotions, Panksepp , developed a neuroscientifically based theory of primary process affects as raw emotional feelings which he associates with evolutionary ingrained subcortical circuits.

Panksepp distinguishes between three distinct kinds of primary process affects, homeostatic, sensory, and emotional. Hence, primary process affect must be somehow assigned to stimuli since otherwise Panksepp could not associate primary process affect with stimuli of such different origins. Analogous to Panksepp, Russell must also presuppose affective assignment though in a slightly different way. He does not associate what he describes as Core Affect itself with a specific type of stimulus since unlike Panksepp he does not speak of sensory, homeostatic, or emotional Core Affect.

Instead, Core Affect is continuously present independent of the presence or absence of particular stimuli. One though has to mention that Russell seems to refer here only to the absence of exteroceptive stimuli since he does not explicitly talk about interoceptive or even neural stimuli in this context.

This means that it cannot be excluded that Core Affect may be related to the assignment of affect to either neural or interoceptive stimuli. Rather than to interoceptive stimuli, Russell explicitly refers to the assignment of affect to exteroceptive stimuli when he describes the transition from Core Affect to emotional episodes and emotional meta-experience.

In the moment when the continuously present Core Affect is related to an episodically occurring exteroceptive stimulus, an emotional episode and meta-experience may occur. This however is possible only if the Core Affect is linked and thus assigned to the exteroceptive stimulus thus presupposing what I here call affective assignment. While both Panksepp and Russell seem to presuppose the assignment of affect to stimuli, the exact functional mechanisms that enable and predispose such affective assignment remain unclear in their accounts.

What functional mechanisms are necessary to enable and predispose the transformation of a non-affective stimulus into an affective one? It is especially worthwhile to consider that the stimulus of interoceptive, exteroceptive, or neural origin must be non-affective. Both Panksepp and Russell consider affect to be essentially subjective rather than objective. Hence, Panksepp distinguishes internal from external and Russell private from public when they characterize Core Affect or primary process affect as subjective rather than objective. One should need to make a conceptual remark here.

The meaning of the term subjective refers here only to the experience of affect, it does not say anything about the underlying neuronal mechanisms that may well be objective. This raises the question how affective assignment makes it possible to transform the originally objective stimulus, interoceptive, exteroceptive, or neural, into a subjective one. Both Russell and Panksepp seem to presuppose some kind of intrinsic stimuli to be crucial in generating affect. Russell does so by explicitly distinguishing Core Affect from extrinsic stimuli and related emotional episodes, while Panksepp argues that neural activity in the subcortical circuits is not dependent upon extrinsic stimuli, i.

This means that both must presuppose some kind of intrinsic activity for the generation of affect Figure 2. The figure shows the relationship between intrinsic activity and stimuli on the left and emotional feeling on the right. The intrinsic activity of the brain interacts with the stimuli that are by themselves non-affective and objective. That rest-stimulus interaction leads to the assignment of affect and subjectivity to the stimulus resulting in emotional feeling.

What could such intrinsic activity be? One may assume it is that activity that can be observed in the absence of any extrinsic stimulation by either intero- or exteroceptive stimuli. Intrinsic means then that the origin of that activity must be traced back to the brain itself as distinguished from body and environment. And it is such resting state activity as intrinsic activity that can be observed in all brain regions cortical and subcortical see Northoff et al. The fact that resting state activity is present throughout the whole brain means that there may already be some neural interactions between the different brain regions within the resting state itself.

For instance the resting state activity level in the sensory cortex may interact with the resting state activity level in the subcortical regions so that one may want to speak of rest—rest interaction. The above described results lend clear empirical support to the assumption that such resting state activity in especially the midline regions is central for emotions and most likely for emotional feelings. And there is further interaction.

Such rest-stimulus interaction may be specified according to the stimulus type either rest-interceptive stimulus interaction or rest-exteroceptive stimulus interaction which in the following I will describe as rest-intero and rest-extero interaction. How do the three types of interaction, rest—rest, rest-intero, and rest-extero relate to affective assignment and more specifically to the non-affective-affective transformation and the objective-subjective transformation?

The resting state activity level is different not only between different regions but even more importantly, different between different persons. This means that the same stimulus encounters a different brain in different persons meaning it must interaction with a different resting state activity level. Due to the individual resting state activity level and its impact on rest-stimulus interaction, the stimulus is processed in a very individual and ultimately private way.

Hence what I called above objective-subjective transformation may correspond on a functional level to rest-stimulus interaction.

Russell and Panksepp may now want to argue that this accounts only for half of the story. Panksepp may say that this leaves emotional affects as based on the stimuli from the BrainMind itself and hence its neural stimuli out; this may be so because rest-stimulus interaction concerns only the interaction with intero- and exteroceptive stimuli.

Hence, my assumption of rest-stimulus interaction corresponding to objective-subjective transformation may well account for what Panksepp calls homeostatic affects and sensory affects, but not emotional affects. Russell may want to make an even stronger point. My assumption of rest-stimulus interaction misses Core Affect altogether because Core Affect is neither related to interoceptive nor exteroceptive stimuli.

Instead the core affect precedes both kind of stimuli that becoming relevant only in emotional episodes. Hence, my assumption that rest-stimulus interaction corresponds to objective-subjective transformation may hold for emotional episodes and emotional meta-experience but not for Core Affect itself. Recent imaging data show that such rest—rest interactions do indeed occur see Northoff et al. In the case of such rest—rest interaction, the resting state activity level of one network is set against that of another network. This has major implications for the conceptual characterization of emotions.

If that very same resting state activity individualizes and privatizes stimuli and their encoding into neural activity, any emotions must be individualized and privatized. That though means that any emotions must necessarily be subjective meaning that it cannot avoid objective-subjective transformation. There is consequently no emotion without emotional feeling with the latter being at the very core of the former. To empirically support this assumption, future studies are needed to test whether the preceding level of resting state activity predicts the degree of especially the subjective-experiential component of emotions, i.

I would hypothesize that the preceding resting state activity predicts especially the subjective-experiential component of emotions. While other components like the vegetative and the cognitive aspects of emotions may rather be predicted by the degree of stimulus-induced activity itself. One may finally raise the question how the here suggested role of the resting state in objective-subjective transformation stands to the above proposed relational coding.

I would argue that both are well compatible. Let me be more specific. The incoming stimulus must be coded in relation to the intrinsic activity level and thus relative to it. The resulting neural activity must then be considered the integral of their interaction, i. That though is possible only if neural activity is coded in terms of a relation between stimulus and intrinsic activity as distinguished from neural coding of the stimulus by itself. I thus assume rest-stimulus and stimulus-rest interaction to presuppose relational coding in very much the same way as the relation between intero- and exteroceptive stimuli is encoded into neural activity see Northoff, a for more details on the question of neural coding.

How about the second feature of affective assignment, that non-affective-affective transformation? What functional mechanisms correspond to the transformation of a non-affective stimulus into an affective one? Panksepp , p. He considers primary process affect to be intrinsically valuative in a wider sense as not being restricted to reward in that it mirrors the value of environmental, bodily, and neural information for the organism.

How can such value be generated, and what kind of functional mechanisms are necessary in order to value stimuli of different origin, exteroceptive, interoceptive, or neural? In order for stimuli of various origins to be valued for the organism, they must be related to the organism itself, including its body and brain. How does affect enter these various interactions? Russell tells us that Core Affect is continuously present even in the absence of exteroceptive stimuli. Unlike exteroceptive stimuli which arise more episodically, there is continuous interoceptive input and thus continuous rest-intero interaction in the brain.

Due to the continuous presence of the body, continuous interoceptive input and subsequent continuous rest-intero interaction cannot be avoided. One may consequently consider rest-intero interaction as one possible candidate functional mechanism that may correspond to what Russell describes as Core Affect. Hence, interaction of the interoceptive stimuli with the neural stimuli may transform the originally non-affective interoceptive stimulus into an affective one. The hypothesis is thus that rest-intero interaction may correspond on the functional level to the non-affective-affective transformation and thus to what Russell described as Core Affect.

However, there is not only Core Affect but also emotional episodes Russell or sensory affect Panksepp in relation to exteroceptive stimuli. How can exteroceptive stimuli be assigned affect and how can they undergo the non-affective-affective transformation?

They may be linked to interoceptive stimuli resulting in an intero-extero interaction. They would thereby be valued, which in turn would lead to a non-affective-affective transformation with the subsequent assignment of affect. Hence, one may consider the interaction of stimuli of various origins with specifically interoceptive stimuli from the body as a necessary condition for the non-affective-affective transformation.

It may also apply to exteroceptive stimuli with intero-extero interaction that may then result in what Russell characterized as emotional episodes and Panksepp as sensory affect. One may now be puzzled. And I considered the interaction of any stimulus with interoceptive stimuli from the body as being necessary for the non-affective-affective transformation. This means that affectivity and subjectivity are co-constituted, which is reflected in both Panksepp and Russell definitions of affect by.

If one interaction takes over at the expense of the respective other, the co-constitution between affectivity and subjectivity may become dysbalanced. This is, for instance, the case in schizophrenia where rest-intero and rest-extero interactions may be reduced leading to an abnormal loss of subjectivity Northoff and Qin, There is thus still non-affective-affective transformation while the objective-subjective transformation seems to fail: These patients thus still experience emotional feelings while their respective contents are no longer experienced as subjective but objective.

While the reverse seems to be the case in depression, where rest-intero interaction seems to predominate over intero-extero interaction Alcaro et al. In the most extreme case, depressed patients say that they no longer feel any emotion, the feeling of non-feeling. Hence, non-affective-affective transformation may be blocked while at the same time this state is experienced as highly subjective implying objective-subjective transformation. The cases of depression and schizophrenia thus indicate the possibility of dissociation between both forms of transformation.

Interoceptive processing and consecutively intero-extero interaction may also be altered or disrupted in alexithymia that concerns the inability to identify and describe emotional feeling. The exteroceptive stimuli and their respective contents can then no longer be associated with emotional feelings: While the contents are experienced as subjective due to functioning rest-intero and rest-extero interactions , they are not assigned emotional feeling due to deficient intero-extero interaction.

Empirically this is supported by a recent study that shows the degree of interoceptive awareness to predict the degree of alexithymia with high interoceptive awareness going along with a low degree of alexithymia Herbert et al. How should emotional feelings be conceptualized on the basis of the intero-exteroceptive relational model of neural coding?

Emotional feelings can no longer be determined in an interoceptive-based way as perceptions of physiological body changes. Instead, emotional feelings may better be described in an neural-intero-exteroceptive relational way thus focusing more on the relation between brain, body and environment than on either the body or the environment itself 3.

This is paradigmatically reflected in what the philosopher Ratcliffe , calls existential feelings.

Emotions & Consciousness (The Secret To True Happiness)

Based on Heidegger, he describes existential feelings as feelings that characterize our relation to the world, i. Most important, emotional feelings like anger, grief, etc. If so, the body itself may only be considered the medium through which feelings can be constituted.

Emotion and consciousness.

Due to the very basic and foundational character of the brain-body-environment relation, the relational concept considers emotional and existential feelings basic and primary for emotions, i. Analogously, the relational view considers our relation to the world primary, basic and crucial to our subjective experience or, as Ratcliffe would probably say, the relation is existential. Since the relational concept characterizes the brain-body-environment as basic, primary, and constitutive of feelings, the here advanced relational concept of emotional feelings seems to complement the empirical approach by Panksepp in conceptual regard.

Feelings and thus affective consciousness can only be primary and basic, as Panksepp claims, because they are our relation to the world. Ellis insistence on the close linkage between motor function and emotional feeling, i. The here suggested relational approach also needs to be distinguished from cognitive approaches. Cognitive approaches focus on the awareness and thus reflection of emotional and cognitive contents thus presupposing access or reflective consciousness, the awareness of subjective experience.

For that various cognitive functions are assumed to be necessary. Since the cognition, i. Future studies may therefore want to investigate how the here described neuronal processes of non-affective-affective and objective-subjective transformation impact cognitive functions and their respective neural substrates.

The relational concept presupposes bilaterally dependent and constitutive linkage between brain, body, and environment. In other terms, motor function must be considered the empirical means by means of which what I conceptually described as relational becomes possible. The figure shows the relationship between emotional feeling and phenomenal consciousness.

This in turn makes possible the generation of emotional feeling and phenomenal consciousness. Following Bennett and Hacker , pp. Either we have pain and subjectively experience or feel pain or we do not feel any pain and then we have no pain. According to Bennett and Hacker , p. By considering feeling as constitutive of emotion and phenomenal consciousness, the relational concept of emotional feeling argues against the explanation of feelings in terms of higher-order cognitive and reflective functions mirroring what is called reflective consciousness.

Phenomenal consciousness, in contrast, does not describe cognitive and behavioral aspects associated with subjective experience. A number of alternative terms and phrases pick out approximately the same core property of phenomenal consciousness. Goldie , pp. How such personal point can be established by relating brain and body to the environment remains to be discussed in detail which however is beyond the scope of this paper see Northoff, , b ; Northoff and Bermpohl, ; Northoff et al.

Finally, the here proposed neuronal mechanisms underlying especially the subjective nature of emotions may be relevant for consciousness in general. The yet to be specified and defined neuronal mechanisms underlying rest-stimulus interaction are assumed to be central for the subjective component.

If so they must be regarded necessary of consciousness in general that is essentially defined as subjective. Even if not sufficient by themselves as neural correlates of consciousness NCC , specific yet unknown ways of rest-stimulus interaction must then be regarded a necessary or predisposing condition of consciousness. I therefore suggested in this paper to complement the embodied concept of emotional feelings by a relational concept that assumes emotional feelings to be constituted by the brain-body-environment relationship.

The relational concept assumes that the environment and the brain itself have not only instrumental and thus indirect impact on emotional feelings via the body but also a direct, e. The present paper focuses on whether such relational concept of emotional feelings is compatible with current empirical data on the neuroscience of emotion processing. If the relational concept of emotional feeling is empirically plausible, even interoceptive awareness should implicate brain regions that process exteroceptive stimuli. This presupposes what I describe as the neural-intero-exteroceptive relational mode of neural coding rather than interoceptive-based translational neural coding see also Northoff, a.

In short I assume relational coding to be the predominant neural code that allows to link emotions to brain, body, and environment. Emotions and emotional feelings may then be considered, metaphorically speaking, the bridge or glue between brain, body, and environment. The intrinsic linkage between brain, body, and environment accounts for the subjective and affective nature of emotional feelings. By being processed in the brain relative to its intrinsic activity at least in the human brain as it is designed emotions cannot avoid becoming subjective and affective.

The subjective and affective components must therefore be regarded intrinsic to and thus defining features of emotions. As such emotions and emotional feeling may be considered paradigmatic cases of consciousness in general which in the current neuroscientific and philosophical debate is rather often neglected see also Northoff, b.

The author declares that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. Is subcortical-cortical midline activity in depression mediated by glutamate and GABA? A cross-species translational approach. Complementary roles of prefrontal cortical regions in cognition, memory, and emotion in primates. Pubmed Abstract Pubmed Full Text. Dead tissue, living ideas: The role of emotion in decision-making: Listening to your heart: Philosophical Foundations of Neuroscience.

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Emotional faces and the default mode network. Craig , , , , , assumes specifically the right insula to be crucially involved which receives autonomic and visceral afferences from lower centers see above and re-represents the interoceptive body state in an integrated way. If these regions mediate interoceptive processing, the question for their role in the subjective experience of bodily and thus interoceptive changes as the basis for emotional feeling arises.

Activity in the right insula also correlated with both the performance in the heartbeat detection task and subjective anxiety symptoms which also correlated with each other. These findings suggest close relationship between interoceptive awareness and emotional feeling. Other studies demonstrated the modulation of these interoceptive stimulus changes by exteroceptive stimuli.

Using fMRI, Critchley , for instance, investigated regional neural activity changes during presentation of happy, sad, angry, and disgusted faces. They observed heart rate changes to be dependent upon the emotional category with sad and angry faces inducing the strongest heart rate changes. According to the authors themselves, these results provide support for the hypothesis that interoceptive stimulus processing may be involved in differentiating between different types of emotional feelings. The group around Pollatos conducted a series of studies on heartbeat perception and emotional feeling.

Using EEG, they distinguished between good and poor heartbeat perceivers. Good heartbeat perceivers Pollatos et al. Taken together, these studies show behaviorally a close relationship between interoceptive awareness, arousal, and emotional feeling.

Significance

The question is whether the above described data support an embodied concept of emotional feeling with exteroceptive stimuli being merely modulatory and instrumental or epiphenomenal. Or whether the data might be interpreted rather in favor of a relational concept of feelings with interoceptive stimuli in relation to exteroceptive stimuli being constitutive and thus central. Presupposing the James—Lange theory, most of the above cited authors have interpreted their data in favor of the interoceptive-based concept. However, I will argue that there are strong arguments which make the data rather compatible with what I call the intero-exteroceptive relational concept of emotional feeling.

I argue that there seems to be a mismatch between empirical data and their interpretation in current imaging studies on emotional feelings and interoceptive processing which I want to support by making the three following points. First, all paradigms employed did not investigate interoceptive stimuli in isolation from exteroceptive stimuli but rather in relation to them. Neural activity changes assumed to be specific for interoceptive awareness thus reflect a relation or dynamic balance between intero- and exteroceptive processing rather than mirroring isolated interoceptive stimulus processing remaining more or less independent of exteroceptive stimulus processing.

Dynamic modulation of the right insula activity as observed by Critchley may thus reflect a dynamic balance between intero- and exteroceptive attention in the heartbeat-auditory tone detection task rather than pure interoceptive heartbeat stimulus processing. Second, neither of the above mentioned studies addressed the question of emotional valence that indicates whether a feeling is positive or negative see also Colombetti, for a discussion of the concept of emotional valence.

Interoceptive awareness may thus be linked to emotional arousal and subjective experience of emotional intensity while it apparently does not seem to determine the valence of the emotional feeling. Regions that have been associated with emotional valence, as distinguished from emotional arousal, include the medial orbitofrontal cortex MOFC , the subgenual and pregenual anterior cingulate cortex PACC , and the ventromedial prefrontal cortex VMPFC; Craig, , ; Phan et al.

The connectivity pattern thus argues strongly in favor of the intero-exteroceptive relational concept of emotional feeling which seems to make isolated interoceptive processing and thus an interoceptive-based concept of emotional feeling rather unlikely. What however is needed to further support this point are investigations of both regional activity and connectivity patterns during intero- and exteroceptive stimulus processing see Hurliman et al.

Third, Pollatos et al. A similar temporal distribution is suggested by Tsuchiya and Adolphs who assume involvement of subcortical regions like brain stem nuclei and hypothalamus that mediate interoceptive stimuli to occur after and later than activation in higher regions like the DMPFC. These higher cortical regions have been associated with processing of higher-order exteroceptive stimuli particularly those that are highly self-related to the organism Northoff and Bermpohl, ; Northoff et al.

The fact that these regions are apparently implicated from early on in interoceptive awareness gives some though indirect support to the assumption that exteroceptive stimuli are involved early in interoceptive processing. Such early involvement indicates that the role of exteroceptive stimulus processing goes beyond mere modulation of interoceptive processing which would be better compatible with late involvement.

In other terms, early involvement of these regions may indicate that interoceptive stimulus processing is coded in relation to exteroceptive stimuli going beyond mere modulation of the former by the latter. The observed early spatio-temporal pattern may thus reflect neural coding of the relationship between intero- and exteroceptive stimulus processing, i.

Otherwise there would be no need for regions predominantly associated with exteroceptive stimulus processing to be implicated so early. While it seems to be less compatible with the assumption of primarily independent interoceptive processing that becomes secondarily modulated by exteroceptive stimuli. Finally, direct empirical support for intero-exteroceptive convergence comes from a recent study by Farb et al. He investigated interoceptive awareness i. While both intero- and exteroceptive awareness yielded dissociable networks i.

Unlike the posterior insula that responded strongly to interoceptive awareness, the anterior insula activity was as much predicted by exteroceptive awareness as interoceptive awareness. Hence, there seems to be intero-exteroceptive convergence in especially the anterior insula with both being integrated in the middle insula as bridge from posterior to anterior parts of the insula. Using biofeedback arousal and relaxation tasks in fMRI, Nagai et al. The level of neural activity in VMPFC and MOFC, which are part of the so-called anterior cortical midline structures aCMS , may thus represent the basal sympathetic or autonomic tone independent of some actual stimuli.

Since the aCMS have been shown to be modulated also by exteroceptive stimuli, neural activity within these regions may mirror a dynamic balance between attention to extero- and interoceptive stimuli see also Nagai et al. This assumption is well compatible with the connectivity pattern of these regions.

The aCMS are also densely connected to regions insula, hypothalamus, and nuclei in the brain stem as such PAG, colliculi, etc. This connectivity pattern predisposes the aCMS for neural processing irrespective of the sensory modality of the respective stimulus, i. The assumption of supramodal processing in aCMS is supported by results from imaging studies. Emotions in either exteroceptive modality visual, auditory, gustatory, olfactory induce neural activity in various regions of the aCMS see above as well as Phan et al.

Finally, stimuli from different origins, i. Taken together, both connectivity pattern and imaging data suggest that neural processing in aCMS is supramodal and domain-independent: Instead it is important how the neural activity in the aCMS is related to the respective intero- or exteroceptive stimulus see below for further discussion. In addition to the aCMS, subcortical midline regions like the periaquaeductal gray PAG , the colliculi, the dorsomedial thalamus, and the ventral striatum may also be considered in processing interoceptive stimuli in relation to exteroceptive ones.

Panksepp ; and also Damasio, , , for instance, assumes that these regions are crucial in constituting emotional feelings. Since the very same regions are also characterized by strong motor connections both afferent and efferent, he and others like Ellis ; unlike Damasio who assumes a sensory-based view of feelings assume emotional feeling to be motor-based. Unfortunately, subcortical regions have often been neglected in imaging studies of emotions which, at least in part, may be due to the fact that neural activity in these regions is rather difficult to reliably visualize in current imaging techniques like fMRI.

However, animal experiments demonstrate the crucial role of these subcortical midline regions in constituting emotional feelings Panksepp, , Future studies in humans are thus needed to investigate subcortical neural activity during emotional feeling in order to bridge the current gap between animals and humans. Furthermore, the relationship between emotional feeling and motor function also needs to be investigated in detail by, for instance, investigating emotional feeling in dependence on variation of motor function and its neural underpinnings and vice versa.

What is the implicit presupposition that drives most of the above cited authors to interpret their data in favor of the James—Lange theory? They seem to presuppose a clear-cut distinction between intero- and exteroceptive stimulus processing with both systems being separate, distinct, and only interacting at specific node points.

According to such view, exteroceptive stimuli are translated into interoceptive stimulus processing whose perception, in turn, is supposed to induce feeling. Exteroceptive stimuli thus have at best an only indirect and mediated impact on emotional feeling in that they must first be translated into interoceptive stimulus processing before they can modulate feelings. I therefore call this model the interoceptive-based translational concept of feeling. However, anatomical connectivity suggests otherwise.

Throughout the brain at all levels both subcortical and cortical and especially in the subcortical-cortical midline system there is convergence between intero- and exteroceptive inputs. This is especially true for regions like the colliculi, the PAG, the tectum, and the aCMS where both intero- and exteroceptive afferences converge onto common neurons see Panksepp, , ; Rolls et al. This suggests that interoceptive stimuli are not only modulated by exteroceptive stimuli at specific node points but rather that the relation, e.

Exteroceptive stimuli are not translated into interoceptive stimulus processing but rather directly and unmediated related to them and it is this relation that seems to be coded in neural activity. The figure compares two different ways of neural coding in emotional feeling. On the left side translational coding describes how intero- and exteroceptive stimuli are separately represented and meta-represented in the neural activity of the brain.

This meta-representation is then perceived which following Damasio and the James—Lange theory leads to emotional feelings. This is different in relational coding on the right side. Here intero- and exteroceptive stimuli are coded in relation to each other with this relation resulting in emotional feeling and subsequent experience of the relationship between body and environment.

Is there any empirical evidence in favor of the intero-exteroceptive relational model of neural coding? Critchley , p. What seems to be coded in the brain is not so much the interoceptive stimulus itself but its relation to the respective exteroceptive stimulus. If neural activity codes the actual relationship and balance between intero- and exteroceptive stimuli, one would expect strong contextual dependence of emotional feelings. The constitution of the emotional feeling, the type of feeling, should then depend on the respective emotional context which implies that different contexts may lead to different types of emotional feelings even in identical situations.

In other terms, the environmental context does not only modulate emotional feelings but actively participates in constituting emotional feelings. If the role of the context were merely modulatory, subjects would not have shown completely different and opposing emotional feelings in the two situations but rather variants of the same feeling. These experiments thus lend further support to the assumption of a constitutive role of the environmental context in emotional feelings rather than remaining merely modulatory.

How are intero- and exteroceptive stimuli related and balanced with each other in relational coding? Rather than coding the intero- or exteroceptive stimulus itself, the degree of correspondence between intero- and exteroceptive stimuli is coded. If, for instance a lion approaches, the heart rate may increase, which may signal strong correspondence and convergence between intero- and exteroceptive stimuli.

This consecutively leads to the constitution of a corresponding emotional feeling, the feeling of fright and anxiety. If, in contrast, the approach of the lion is not accompanied by heart rate increases, as for instance if one is not clear whether the lion is real or not, there may be a mismatch between intero- and exteroceptive stimuli. This may result in a different emotional feeling, the feeling of doubt and hesitation. The degree of convergence and divergence between intero- and exteroceptive stimuli may thus determine the kind of emotional feeling.

That is well in accordance with the relational concept rather than with the translational one that claims for an interoceptive- and thus bodily based approach. Interoceptive stimulus processing remaining isolated, unrelated and independent from exteroceptive stimulus processing is consequently assumed to remain principally impossible.

This implies what I call intero-exteroceptive relational coding while it excludes interoceptive-based translational coding. What does this imply in experimental regard? The experimental efforts to isolate interoceptive stimulus processing and to search for its specific neural correlates may be futile since exteroceptive stimulus processing may always already be implicated in interoceptive stimulus processing. The philosopher Hurley , pp. If the relationship between input and output is indirect and thus merely instrumental, changes in perceptual content are dependent upon changes in the input.

Every change in motor output has to modulate sensory input in order to have an impact on perceptual content implying that the output can not change independently of the input: What does this mean with regard to emotional feelings and their relation to the environment? Presupposing instrumental dependence, the environment can impact emotional feelings only indirectly via the body, i. This is nicely reflected in a quote from a recent paper about emotion and consciousness: Non-instrumental dependence, in contrast, is described by Hurley as direct dependence of perceptual content on motor output independent of sensory input; even if the sensory input remains the same and fixed, perceptual content can vary depending on motor output.

This means that motor output has direct access to perceptual content independent of sensory input and therefore no longer operates indirectly via sensory input as in instrumental dependence; instead, perceptual content may vary in orientation on motor output independent of sensory input and thus directly. What does such non-instrumental or constitutional, as I will call it in the following see also Colombetti, , dependence imply for the relationship between body and environment in emotional feeling? If the relationship between emotional feeling and environment is direct and therefore constitutional, i.

The environment itself may then directly involved in constituting emotional feelings. Thereby, the concept of environment is meant here in a social sense, social environment, as distinguished from the merely physical world or physical environment. This has empirically been paradigmatically exemplified in a recent study on reward Fliessbach et al. Two subjects a and b were simultaneously scanned while receiving rewards. While the reward for the subject a was fixed, the one for subject b was varied; this and the converse case, increasing rewards for subject a and fixed rewards for subject b, allowed to investigate its impact of the environment, i.

Interestingly, emotional feelings and neural activity in reward circuitry in subject a did not so much depend on the size of the reward it received but rather on the relation of or balance to its own reward when compared to the one received by subject b. It is such constitutional, i. Such intrinsic linkage between emotional feelings and the social environment is empirically further supported by the observed overlap between emotion processing and social processing like social intentions; see Ciaramidaro et al.

These and other data end strong support to an intrinsically social and thus relational concept of emotional feeling. The relational approach shifts the focus of attention from the body, as in the embodied approach, to the role of the environment in emotional feelings.

Rather than modulating emotional feelings indirectly via bodily representations, the environment is supposed to be involved directly in constituting emotional feelings. How does the person-environment relation account for the variety of different specific emotional feelings? The lack of specificity concerning distinct emotions has often been criticized in feeling theories like the James—Lange theory see also Niedenthal et al. Autonomic bodily changes like arousal are rather unspecific reactions that do not allow to distinguish between distinct emotions. This criticism has been furnished by the Schachter and Singer experiments demonstrating that subjects with autonomous nervous system stimulation, as induced by epinephrine, experienced the resulting arousal as either anger or euphoria in dependence on the respective context they were placed in a room with either an angry or happy actor.

The conclusion is often drawn that physiological bodily changes and arousal themselves remain unspecific and cannot contribute to determine specific emotions ; determination and distinction of specific feelings can consequently not be based upon physiological bodily changes but must be found elsewhere. This argument of the lack of specificity of bodily representations has been countered in different ways by referring to motor, cognitive, or neural representation.

Zajonc , , for instance, claims that the motor system allows for extremely subtle distinctions which means that even a number of limited bodily states can support a very large number of representational distinctions of distinct emotional feelings. Rather than referring to motor capacities, cognitive theories, e. Damasio , , suggested a middle way between motor and cognitive representation. He focuses on those subcortical brain regions that register physiological bodily states which may allow for a wider representational spectrum than the muscles and viscera themselves that are actually represented in the respective neural states.

All these approaches have in common that they still presuppose representation of emotional feelings in motor, cognitive, or neural-subcortical functions. The relational approach, in contrast, claims that the wide variety of different emotional feelings may ultimately be traced back to the relation between person and environment rather than to motor, cognitive, or neural-subcortical representation.

Since an abundant variety of different person-environment relations are possible, different emotional feelings can be constituted. The question for the specificity of emotional feelings is thus traced back to the possible and impossible person-environment relations rather than to the representational capacities of specific functions, i.

If emotional feelings are intrinsically relational, i. What Ratcliffe calls existential feeling presupposes what I here advance as relational concept of emotional feeling. How does the person and thus the subject come into play in emotional feelings? This will be the focus in the next sections. I so far focused on the relationship between intero- and exteroceptive stimulus processing in emotional feeling. Neurobiologically this was supposed to be related to the anatomical convergence between the respective pathways and the kind of coding, i.

While conceptually this implied a shift from the embodied concept of emotional feeling to a relational concept or as the philosophers may want to say an extended concept. This pointed out the intrinsic relation to emotional feelings to the environment as bridge between body and environment. What remains unclear though are two aspects: And why are emotional feelings affective and thus emotional? Taking a purely logical stance one could well imagine mere feelings without any emotions.

Both questions dent deeply in various domains of research including consciousness see Northoff, a , b which though I will avoid here to keep matters simple. I will here focus only on some neurobiological mechanisms while leaving the philosophical implications open. In order to shed some light on these questions, I turn to recent results about the relation between resting state activity and emotions.

A recent study by Sreenivas et al.

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From Emotions to Consciousness – A Neuro-Phenomenal and Neuro-Relational Approach.
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They demonstrated that sad faces induced a higher degree of deactivation, i. In contrast, activation and thus positive emotional signal changes were observed in the lateral fronto-parietal regions except in left middle frontal gyrus. Finally, functional connectivity pattern also differed between sad and happy emotions for the connections between the midline and the lateral regions with VMPFC, PCC, and precuneus being central nodes. While this study demonstrates that emotions are associated with midline regions that show high resting state activity, it leaves open whether that is related to intero- or exteroceptive stimuli.

This was tested for by a study by Wiebking et al. These served to subtract both intero- and exteroceptive signal changes from the resting state which, as expected, yielded higher activity changes in the midline regions. We then determined the degree of deactivation during both intero- and exteroceptive awareness. That yielded stronger deactivation in exteroceptive awareness when compared to interoceptive awareness. How is all that related to emotions? For that Wiebking et al. Interestingly, we did not observe any correlation of the emotion measures with the midline signal changes during interoceptive awareness alone.

The stronger the emotion score, the smaller the degree of deactivation in the midline regions thus being closer to the resting state activity level. In contrast, no correlation was observed with signal changes during interoceptive awareness. Hence, these results underline the central importance of intrinsic and thus resting state activity for emotions. These results show the strong association between resting state activity and emotions.

They though leave open whether emotions can also modulate resting state activity or whether the latter predict the former. Several recent studies demonstrated the prediction of stimulus-induced activity by the preceding resting state activity implying rest-stimulus interaction see Northoff et al. This was mainly shown in the in the sensory domain while, as to my knowledge, such studies are not yet available in the domain of emotions. There are though a couple of studies that show the reverse, modulation of resting state activity by preceding emotions.

Focusing on emotions, Eryilmaz et al. They asked the participants after the resting state period about their thoughts. Neuronally, they showed that the resting state periods after fearful faces showed higher neuronal activity in subcortical regions pallidum, anterior thalamus, hypothalamus than the ones following neutral movies rest after fearful larger than rest after neutral.

This means that the inclusion of fearful emotions in the preceding movie had a clear effect on the level of subsequent resting state activity. This study clearly demonstrates that emotions have an impact on the subsequent resting state thus implying what we coined as stimulus-rest interaction see Northoff et al. This was also observed in another study. This revealed increased functional connectivity from the amygdala to the cortical midline structures like the MPFC, the PCC, and the precuneus. This indicates that psychological stress implicating strong and negative emotions can affect the subsequent resting state activity thus implying stimulus-rest interaction.

The close relationship between emotions and resting state is further supported albeit indirectly by the observation of severe resting state alterations in major depressive disorder see Alcaro et al. For that I turn to two of the major proponents of emotional feeling, Jaak Panksepp and Jim Russell, and how they conceptualize especially the subjective-experiential component of emotional feeling. This will be then linked in subsequent sections to the above described findings of the close relationship between intrinsic activity and emotions.

Based on the centrality of affect and emotions, Panksepp , developed a neuroscientifically based theory of primary process affects as raw emotional feelings which he associates with evolutionary ingrained subcortical circuits. Panksepp distinguishes between three distinct kinds of primary process affects, homeostatic, sensory, and emotional. Hence, primary process affect must be somehow assigned to stimuli since otherwise Panksepp could not associate primary process affect with stimuli of such different origins.

Analogous to Panksepp, Russell must also presuppose affective assignment though in a slightly different way. He does not associate what he describes as Core Affect itself with a specific type of stimulus since unlike Panksepp he does not speak of sensory, homeostatic, or emotional Core Affect. Instead, Core Affect is continuously present independent of the presence or absence of particular stimuli.

One though has to mention that Russell seems to refer here only to the absence of exteroceptive stimuli since he does not explicitly talk about interoceptive or even neural stimuli in this context. This means that it cannot be excluded that Core Affect may be related to the assignment of affect to either neural or interoceptive stimuli.

Rather than to interoceptive stimuli, Russell explicitly refers to the assignment of affect to exteroceptive stimuli when he describes the transition from Core Affect to emotional episodes and emotional meta-experience. In the moment when the continuously present Core Affect is related to an episodically occurring exteroceptive stimulus, an emotional episode and meta-experience may occur.

This however is possible only if the Core Affect is linked and thus assigned to the exteroceptive stimulus thus presupposing what I here call affective assignment. While both Panksepp and Russell seem to presuppose the assignment of affect to stimuli, the exact functional mechanisms that enable and predispose such affective assignment remain unclear in their accounts. What functional mechanisms are necessary to enable and predispose the transformation of a non-affective stimulus into an affective one? It is especially worthwhile to consider that the stimulus of interoceptive, exteroceptive, or neural origin must be non-affective.

Both Panksepp and Russell consider affect to be essentially subjective rather than objective. Hence, Panksepp distinguishes internal from external and Russell private from public when they characterize Core Affect or primary process affect as subjective rather than objective.

Supplemental Content

The James–Lange theory considers emotional feelings as perceptions of physiological body changes. The relational approach thus goes together with what may be described as neuro-phenomenal approach. Such neuro-phenomenal approach does not only inform emotions and emotional feeling. Trends Cogn Sci. Apr;11(4) Epub Feb Emotion and consciousness. Tsuchiya N(1), Adolphs R. Author information: (1)California Institute.

One should need to make a conceptual remark here. The meaning of the term subjective refers here only to the experience of affect, it does not say anything about the underlying neuronal mechanisms that may well be objective. This raises the question how affective assignment makes it possible to transform the originally objective stimulus, interoceptive, exteroceptive, or neural, into a subjective one. Both Russell and Panksepp seem to presuppose some kind of intrinsic stimuli to be crucial in generating affect. Russell does so by explicitly distinguishing Core Affect from extrinsic stimuli and related emotional episodes, while Panksepp argues that neural activity in the subcortical circuits is not dependent upon extrinsic stimuli, i.

The figure shows the relationship between intrinsic activity and stimuli on the left and emotional feeling on the right. The intrinsic activity of the brain interacts with the stimuli that are by themselves non-affective and objective. That rest-stimulus interaction leads to the assignment of affect and subjectivity to the stimulus resulting in emotional feeling. What could such intrinsic activity be?

One may assume it is that activity that can be observed in the absence of any extrinsic stimulation by either intero- or exteroceptive stimuli. Intrinsic means then that the origin of that activity must be traced back to the brain itself as distinguished from body and environment. And it is such resting state activity as intrinsic activity that can be observed in all brain regions cortical and subcortical see Northoff et al. The fact that resting state activity is present throughout the whole brain means that there may already be some neural interactions between the different brain regions within the resting state itself.

And there is further interaction. Such rest-stimulus interaction may be specified according to the stimulus type either rest-interceptive stimulus interaction or rest-exteroceptive stimulus interaction which in the following I will describe as rest-intero and rest-extero interaction. How do the three types of interaction, rest—rest, rest-intero, and rest-extero relate to affective assignment and more specifically to the non-affective-affective transformation and the objective-subjective transformation?

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My assumption of rest-stimulus interaction misses Core Affect altogether because Core Affect is neither related to interoceptive nor exteroceptive stimuli. Instead the core affect precedes both kind of stimuli that becoming relevant only in emotional episodes. Hence, my assumption that rest-stimulus interaction corresponds to objective-subjective transformation may hold for emotional episodes and emotional meta-experience but not for Core Affect itself.

Recent imaging data show that such rest—rest interactions do indeed occur see Northoff et al. In the case of such rest—rest interaction, the resting state activity level of one network is set against that of another network. This has major implications for the conceptual characterization of emotions. If that very same resting state activity individualizes and privatizes stimuli and their encoding into neural activity, any emotions must be individualized and privatized.

That though means that any emotions must necessarily be subjective meaning that it cannot avoid objective-subjective transformation. There is consequently no emotion without emotional feeling with the latter being at the very core of the former. To empirically support this assumption, future studies are needed to test whether the preceding level of resting state activity predicts the degree of especially the subjective-experiential component of emotions, i. I would hypothesize that the preceding resting state activity predicts especially the subjective-experiential component of emotions.

While other components like the vegetative and the cognitive aspects of emotions may rather be predicted by the degree of stimulus-induced activity itself. One may finally raise the question how the here suggested role of the resting state in objective-subjective transformation stands to the above proposed relational coding.